Ecological effects of reduced nutrient loading (oligotrophication) on lakes: an introduction

1. The variable ecological response of lakes to reduced nutrient loading (oligotrophication) at sites in Europe and North America was discussed at a workshop held in Silkeborg (Denmark) in January 2003. Studies of lake oligotrophication were presented based on both long‐term monitoring and data generated by palaeolimnological methods. 2. This introduction to the special issue provides short summaries of a series of the papers presented and their limnological context. Results show that the majority of lakes had approached a new equilibrium of phosphorus (P) and nitrogen (N) concentrations 10–15 years (P) and 0–5 years (N) after a major reduction in loading, irrespective of hydraulic retention time. Phytoplankton biomass decreased and a shift towards meso‐oligotrophic species dominance occurred. The fish responded surprisingly fast to the loading reduction in most lakes. As a result, the percentage of piscivores increased and total fish biomass declined markedly, which may explain an increase in the body size of cladocerans and an increase in the zooplankton to phytoplankton biomass ratio seen in many of the lakes. 3. Monitoring has in general been initiated after the effects of eutrophication became apparent. In this context palaeolimnological techniques become very useful because they allow limnologists to extend time scales of coverage and to define restoration targets and baseline conditions. Moreover, lake sediments pre‐dating anthropogenic disturbance can be used to examine ecological response to, for instance, climate variability, allowing problems associated with multiple stressors to be addressed. 4. It is concluded that there is a great need for a synthetic, holistic approach to studying lake oligotrophication, combining multiple techniques of palaeolimnological sediment analysis with detailed but temporally limited long‐term monitoring of chemical and biological variables. This is important, not least to assess future responses to nutrient loading reductions, as global warming will interact with a range of external stressors and ultimately affect lake management strategies to deal with the resultant feedbacks.     

Stomatal crypts may facilitate diffusion of CO2 to adaxial mesophyll cells in thick sclerophylls

In some plants, stomata are exclusively located in epidermal depressions called crypts. It has been argued that crypts function to reduce transpiration; however, the occurrence of crypts in species from both arid and wet environments suggests that crypts may play another role. The genus Banksia was chosen to examine quantitative relationships between crypt morphology and leaf structural and physiological traits to gain insight into the functional significance of crypts. Crypt resistance to water vapour and CO₂ diffusion was calculated by treating crypts as an additional boundary layer partially covering one leaf surface. Gas exchange measurements of polypropylene meshes confirmed the validity of this approach. Stomatal resistance was calculated as leaf resistance minus calculated crypt resistance. Stomata contributed significantly more than crypts to leaf resistance. Crypt depth increased and accounted for an increasing proportion of leaf resistance in species with greater leaf thickness and leaf dry mass per area. All Banksia species examined with leaves thicker than 0.6 mm had their stomata in deep crypts. We propose that crypts function to facilitate CO₂ diffusion from the abaxial surface to adaxial palisade cells in thick leaves. This and other possible functions of stomatal crypts, including a role in water use, are discussed.     

High-frequency field measurements of diurnal carbon isotope discrimination and internal conductance in a semi-arid species, Juniperus monosperma

We present field observations of carbon isotope discrimination (Δ) and internal conductance of CO₂ ( g _i) collected using tunable diode laser spectroscopy (TDL). Δ ranged from 12.0 to 27.4‰ over diurnal periods with daily means from 16.3 ± 0.2‰ during drought to 19.0 ± 0.5‰ during monsoon conditions. We observed a large range in g _i, with most estimates between 0.04 and 4.0  µ mol m⁻² s⁻¹ Pa⁻¹. We tested the comprehensive Farquhar, O'Leary and Berry model of Δ (Δ_comp), a simplified form of Δ_comp (Δ_simple) and a recently suggested amendment (Δ_revised). Sensitivity analyses demonstrated that varying g _i had a substantial effect on Δ_comp, resulting in mean differences between observed Δ (Δ_obs) and Δ_comp ranging from 0.04 to 9.6‰. First-order regressions adequately described the relationship between Δ and the ratio of substomatal to atmospheric CO₂ partial pressure ( p _i/ p _a) on all 3 d, but second-order models better described the relationship in July and August. The three tested models each best predicted Δ_obs on different days. In June, Δ_simple outperformed Δ_comp and Δ_revised, but incorporating g _i and all non-photosynthetic fractionations improved model predictions in July and August.     

A NEW TRISTICHOPTERID (SARCOPTERYGII, TETRAPODOMORPHA) FROM THE UPPER FAMENNIAN EVIEUX FORMATION (UPPER DEVONIAN) OF BELGIUM

Abstract:  Additional material of a large specimen of tristichopterid fish from the Upper Famennian Evieux Formation of Belgium is described. This large fish was previously assigned to Tristichopteridae gen. et sp. indet. due to the lack of diagnostic anatomical data. New available material consists of the internal surface of the parietal shield, vomers and anterior part of the parasphenoid, subopercular and submandibulo-branchiostegal bones, and an internal view of the anterior part of the mandible. A possible autapomorphy of the new form from Belgium, Langlieria socqueti gen. nov. et sp. nov., is the absence of marginal teeth on the vomer except on its most lateral part. Apart from these features, it only differs from the genus Mandageria from Australia in the absence of marginal teeth between the dentary fang and the mandibular symphysis, in the presence of a raised marginal crest lateral to the anterior coronoid fang, and in the presence of numerous small marginal teeth on the premaxilla. It differs from the cosmopolitan genus Eusthenodon in a number of respects: the supratemporal, tabular, and postparietal bones are superficially fused, as are the intertemporal and parietal bones, the dermal ornament is proportionally very fine, and the denticulated field of the parasphenoid stands proud rather than being recessed into the body of the bone.     

Dynamic micro-geographic and temporal genetic diversity in vertebrates: the case of lake-spawning populations of brown trout (Salmo trutta)

Conservation of species should be based on knowledge of effective population sizes and understanding of how breeding tactics and selection of recruitment habitats lead to genetic structuring. In the stream‐spawning and genetically diverse brown trout, spawning and rearing areas may be restricted source habitats. Spatio–temporal genetic variability patterns were studied in brown trout occupying three lakes characterized by restricted stream habitat but high recruitment levels. This suggested non‐typical lake‐spawning, potentially representing additional spatio–temporal genetic variation in continuous habitats. Three years of sampling documented presence of young‐of‐the‐year cohorts in littoral lake areas with groundwater inflow, confirming lake‐spawning trout in all three lakes. Nine microsatellite markers assayed across 901 young‐of‐the‐year individuals indicated overall substantial genetic differentiation in space and time. Nested gene diversity analyses revealed highly significant (≤P = 0.002) differentiation on all hierarchical levels, represented by regional lakes (F_LT = 0.281), stream vs. lake habitat within regional lakes (F_HL = 0.045), sample site within habitats (F_SH = 0.010), and cohorts within sample sites (F_CS = 0.016). Genetic structuring was, however, different among lakes. It was more pronounced in a natural lake, which exhibited temporally stable structuring both between two lake‐spawning populations and between lake‐ and stream spawners. Hence, it is demonstrated that lake‐spawning brown trout form genetically distinct populations and may significantly contribute to genetic diversity. In another lake, differentiation was substantial between stream‐ and lake‐spawning populations but not within habitat. In the third lake, there was less apparent spatial or temporal genetic structuring. Calculation of effective population sizes suggested small spawning populations in general, both within streams and lakes, and indicates that the presence of lake‐spawning populations tended to reduce genetic drift in the total (meta‐) population of the lake.     

Restoring lakes by using artificial plant beds: habitat selection of zooplankton in a clear and a turbid shallow lake

1. Return of large‐bodied zooplankton populations is of key importance for creating a shift from a turbid to a clear‐water state in shallow lakes after a nutrient loading reduction. In temperate lakes, recovery is promoted by submerged macrophytes which function as a daytime refuge for large zooplankton. However, recovery of macrophytes is often delayed and use of artificial plant beds (APB) has been suggested as a tool to enhance zooplankton refuges, thereby reinforcing the shift to a clear‐water state and, eventually, colonisation of natural plants. 2. To further evaluate the potential of APB in lake restoration, we followed the day–night habitat choices of zooplankton throughout summer in a clear and a turbid lake. Observations were made in the pelagic and littoral zones and in APB in the littoral representing three different plant densities (coverage 0%, 40% and 80%). 3. In the clear lake, the zooplankton (primarily Daphnia) were mainly found in the pelagic area in spring, but from mid‐May they were particularly abundant in the APB and almost exclusively so in mid‐June and July, where they appeared in extremely high densities during day (up to 2600 ind. L⁻¹). During night Daphnia densities were overall more equally distributed between the five habitats. Ceriodaphnia was proportionally more abundant in the APB during most of the season. Cyclopoids were more abundant in the high APB during day but were equally distributed between the five habitats during night. 4. In the turbid lake, however, no clear aggregation was observed in the APB for either of the pelagic genera (Daphnia and Bosmina). This may reflect a higher refuge effect in the open water due to the higher turbidity, reduced ability to orient to plant beds and a significantly higher fish density (mainly of roach, Rutilus rutilus, and perch, Perca fluviatilis) in the plant beds than in the clear lake. Chydorus was found in much higher proportions among the plants, while cyclopoids, particularly the pelagic Cyclops vicinus, dominated in the pelagic during day and in the pelagic and high density plants during night. 5. Our results suggest that water clarity is decisive for the habitat choice of large‐bodied zooplankton and that introduction of APB as a restoration measure to enhance zooplankton survival is only a useful tool when water clarity increases following loading reduction. Our results indicate that dense APB will be the most efficient.     

Is biofuel policy harming biodiversity in Europe?

We assessed the potential impacts of land-use changes resulting from a change in the current biofuel policy on biodiversity in Europe. We evaluated the possible impact of both arable and woody biofuel crops on changes in distribution of 313 species pertaining to different taxonomic groups. Using species-specific information on habitat suitability as well as land use simulations for three different biofuel policy options, we downscaled available species distribution data from the original resolution of 50 to 1 km. The downscaled maps were then applied to analyse potential changes in habitat size and species composition at different spatial levels. Our results indicate that more species might suffer from habitat losses rather than benefit from a doubled biofuel target, while abolishing the biofuel target would mainly have positive effects. However, the possible impacts vary spatially and depend on the biofuel crop choice, with woody crops being less detrimental than arable crops. Our results give an indication for policy and decision makers of what might happen to biodiversity under a changed biofuel policy in the European Union. The presented approach is considered to be innovative as to date no comparable policy impact assessment has been applied to such a large set of key species at the European scale.     

Reconstruction of the pharyngeal corpus of Aphelenchus avenae (Nematoda: Tylenchomorpha), with implications for phylogenetic congruence

The corpus of the pharynx in the nematode Aphelenchus avenae (Nematoda: Tylenchomorpha) was three‐dimensionally reconstructed to address questions of phylogenetic significance. Reconstructed models are based on serial thin sections imaged by transmission electron microscopy. The corpus comprises six classes of radial cells, two classes of marginal cells, and 13 neurones belonging to eight classes. Between the arcade syncytia and isthmus cells, numbers of cell classes along the pharyngeal lumen and numbers of nuclei per cell class correspond exactly between A. avenae and Caenorhabditis elegans. The number of radial cell classes between the arcade syncytia and the dorsal gland orifice (DGO) in A. avenae is also identical with outgroups. Proposed homologies of the pharynx imply that expression of the anterior two cell classes as epithelial or muscular differs within both Rhabditida and Tylenchomorpha. Numbers of neurone cell bodies within the corpus correspond exactly to C. elegans, other free‐living outgroups, and other Tylenchomorpha. Neurone polarity and morphology support conserved relative positions of cell bodies of putative neurone homologues. The configuration of cells in the procorpus, including the length of individual cell classes along its lumen, differs across representatives of three deep Tylenchomorpha lineages. Nonhomology of the procorpus challenges the homology of DGO position within the metacorpus, the primary taxonomic character for circumscribing ‘Aphelenchoidea’. Comparison of A. avenae with Aphelenchoides blastophthorus shows that, despite gross pharynx similarity, these nematodes have several differences in corpus construction at a cellular level. The possibility of convergent evolution of an ‘aphelenchid’ pharynx in two separate lineages would be congruent with molecular‐based phylogeny. Putative homologies and conserved arrangement of pharyngeal neurones in Tylenchomorpha expand the experimental model of C. elegans. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010.     

Systematic significance of tepal surface morphology in tribes Persicarieae and Polygoneae (Polygonaceae)

In order to improve understanding of the systematic relationships between taxa and of tepal-pollinator and tepal-habitat relationships, a detailed scanning electron microscopy (SEW study of the adaxial tepal surface of 135 taxa belonging to c. 14 genera in the tribes Persicarieae and Polygoneae (Polygonoideae-Polygonaceae) was carried out. The tepal surfaces have revealed some striking differences among the genera studied. Three main patterns of sculpturing and cell shape can be recognized. Type I: mostly rectangular to elongate cells with straight or undulating anticlinal walls; the cuticles smooth or striate in longitudinal direction and often continuous (Persicaria, Bistortu, Aconogonon, Koenigia and some species of Fagopyrum). Type II: irregular tesselated to elongated cells, rarely rectangular with mostly sinuate anticlinal walls; the cuticles rarely with longitudinal striation, but with strong orthogonal to reticulate ridges or striae, often without correlation between cells (Polygonum s.s., Polygonella, Fallopia, Parapteropyrum, Pteropyrum, Atraphaxis and Calligonum). Type III: mostly irregular, mammilliform, papillose cells with either longitudinal tightly pressed or randomly ridged cuticles (some species of Fagotyrum and Oxygonum). The present data generally support the division of Polygoneae and Persicarieae by Ronse Decraene & Akeroyd. The mammilliform, papillate epidermis of tepals in Oxygonum and some Fagopyrum is probably well-adapted to entomophily either as a foothold or even as a light-trap, and also shows some correlation with heterostyly. It is noteworthy that the tepal structure of Polygonella (tesselated form) is highly similar to that of Polygonum s. s. (especially P. sect. Duravia). Cuticular striation alone does not appear to have much systematic value, but has a mechanical function in affording extra strength to the cell walls. Some correlations between the degree/shape of cuticles and the pollination mechanisms or the habitat of each taxon are also briefly discussed. It is concluded that the studied tepal characters can be useful in improving the classification of tribes Persicarieae and Polygoneae, arid give additional support at the infrageneric level.